TREATISE ONLINE - College of William and Mary

TREATISE 

ONLINE 

Number 29 

Part N, Revised, Volume 1, Chapter 24: 

Extinction in the Marine Bivalvia 

Paul G. Harnik and Rowan Lockwood 

2011 

Lawrence, Kansas, USA 

ISSN 2153-4012 (online) 

paleo.ku.edu/treatiseonline

PART N, REVISED, VOLUME 1, CHAPTER 24: 

EXTINCTION IN THE MARINE BIVALVIA 

Paul G. Harnik and rowan lockwood 

[Department of Geological and Environmental Sciences, Stanford University, paulharnik@gmail.com; and Department of Geology, The College of 

William and Mary, rxlock@wm.edu] 

Bivalves are diverse and abundant constituents 

of modern marine faunas, and they 

have a rich fossil record that spans the 

Phanerozoic (Hallam & miller, 1988; 

mcroberts, 2001; Fraiser & bottjer, 

2007). Due to the high quality of their 

fossil record, they are well suited for investigating 

patterns of biodiversity change and 

the processes that generate these patterns. 

Extinction and its influence on patterns of 

diversification in the Bivalvia have figured 

prominently in a number of previous studies. 

Did bivalves diversify exponentially over 

geologic time with little long-term influence 

of mass extinction events (stanley, 

1975, 1977), or were catastrophic extinction 

events critical in shaping their dramatic post- 

Paleozoic radiation (Gould & calloway, 

1980)? Extinction is an important process 

in the evolutionary history of many clades. 

Selective or chance survivorship can shape 

morphological, ecological, and phylogenetic 

diversity and disparity (roy & Foote, 1997; 

jablonski, 2005; erwin, 2008). Extinction 

selectivity can also affect the susceptibility 

of lineages to later periods of environmental 

change (stanley, 1990a; jackson, 1995; 

jablonski, 2001; roy, Hunt, & jablonski, 

2009). In addition, extinction can open up 

opportunities for diversification through 

the removal of incumbent taxa (walker & 

Valentine, 1984; rosenzweiG & mccord, 

1991; bambacH, knoll, & sePkoski, 2002; 

jablonski, 2008b). Understanding how 

and why some organisms, including many 

bivalves, and not others became extinct in 

the past may prove useful in predicting the 

response of modern marine ecosystems to 

environmental change (dietl & Flessa, 

2009). 

Here we briefly review those features 

of the bivalve fossil record that make it 

particularly suitable for investigating diversity 

dynamics over geologic time. We then 

introduce recently developed analytical 

methods for estimating rates of extinction 

and origination from paleontological data 

that account for temporal variation in the 

quality of the preserved and sampled fossil 

record. Applying these methods to data for 

marine bivalves, we present a new analysis 

of extinction, origination, and preservation 

rates for bivalve genera over the Phanerozoic 

and examine the effect of extinction rate on 

subsequent origination rate. We review the 

growing literature on extinction risk in fossil 

marine bivalves and summarize the roles of 

several biological factors that have proven 

important in predicting survivorship over 

geologic time. Although recent and historical 

extinction in freshwater mussels has been 

well studied (williams & others, 1993; 

ricciardi & rasmussen, 1999; lydeard 

& others, 2004; strayer & others, 2004; 

boGan, 2006), we focus on marine bivalves 

due to the quality of their fossil record over 

long time scales and the general congruence 

between phylogenetic hypotheses and 

morphologic taxonomies (jablonski & 

Finarelli, 2009). 

MARINE BIVALVES AS 

A MODEL SYSTEM FOR 

ECOLOGICAL AND 

EVOLUTIONARY ANALYSIS 

The marine bivalve fossil record has been 

studied intensively by paleontologists and 

malacologists for centuries. This body of 

work has produced a detailed picture of the 

history of the clade and has advanced our 

general understanding of the processes that 

generate and maintain biodiversity in marine 

systems over time. Data for fossil bivalves 

© 2011, The University of Kansas, Paleontological Institute, ISSN (online) 2153-4012 

Harnik, Paul G., & Rowan Lockwood. 2011. Part N, Revised, Volume 1, Chapter 24: Extinction 

in the marine Bivalvia. Treatise Online 29:1–24, 4 fig., 1 table, 1 appendix.

2 Treatise Online, number 29 

have been instrumental in informing debates 

concerning the roles of biological factors in 

extinction risk (e.g., stanley, 1986a; rauP 

& jablonski, 1993; jablonski, 2005; riVadeneira 

& marquet, 2007; cramPton & 

others, 2010), the tempo and mode of evolutionary 

change (e.g., kelley, 1983; Geary, 

1987; rooPnarine, 1995), the processes 

underlying geographic gradients in diversity 

(e.g., roy & others, 1998; crame, 2002; 

Vermeij, 2005; jablonski, roy, & Valentine, 

2006), and the role of predation in 

evolutionary trends (e.g., stanley, 1986b; 

kelley, 1989; dietl & others, 2002). This 

depth of study is due in part to the high 

preservation potential of bivalve shells in 

shallow marine environments. 

Marine bivalves are not free from the 

taphonomic insults experienced by other 

marine invertebrate taxa, but their fossil 

record is relatively complete (Valentine, 

1989; Foote & rauP, 1996; HarPer, 1998; 

Foote & sePkoski, 1999; kidwell, 2005; 

Valentine & others, 2006), and the taphonomic 

biases that affect this record are increasingly 

well understood (cooPer & others, 

2006; Valentine & others, 2006). Taxa that 

have readily soluble shell microstructures, are 

small-bodied or thin-shelled, geographically 

restricted, commensal or parasitic, epifaunal, 

and/or occur in deeper water are less likely to 

be preserved and sampled (cooPer & others, 

2006; Valentine & others, 2006). Yet, the 

probability of being preserved and sampled 

is relatively high for bivalves living at shelf 

to intertidal depths. Approximately 75% of 

all living genera and subgenera of shallow 

marine bivalves are also known from the fossil 

record (Valentine & others, 2006). Although 

postmortem dissolution of primary shell 

aragonite has resulted in considerable loss 

of molluscan skeletal material from the rock 

record (cHerns & wriGHt, 2000, 2009), this 

taphonomic filter does not appear to have 

biased macroevolutionary patterns inferred 

from fossil mollusks (kidwell, 2005). 

Recent studies have shown significant 

agreement between ecological metrics calculated 

for molluscan death assemblages and 

the living communities from which they 

are derived (kidwell, 2001, 2002, 2005; 

lockwood & cHastant, 2006; Valentine 

& others, 2006). Notably, instances 

in which the ecological agreement between 

life and death assemblages is poor tend to 

be associated with sites affected by recent 

and pronounced anthropogenic environmental 

change (e.g., eutrophication and 

benthic trawling), and not postmortem shell 

loss (kidwell, 2007). These taphonomic 

analyses provide a foundation for examining 

ecological shifts in the Bivalvia over geologic 

time as well as the susceptibility of bivalve 

taxa with particular traits to extinction. 

ESTIMATING EXTINCTION 

AND ORIGINATION FROM 

INCOMPLETE DATA 

Accurately estimating extinction and origination 

rates is challenging for all taxa, due to 

incomplete observations. In paleontological 

studies, the observed stratigraphic distribution 

of fossil occurrences is affected by preservation 

and sampling, leading to temporal 

offsets between a taxon’s true time of origination 

and extinction and its observed first 

and last occurrences (siGnor & liPPs, 1982; 

marsHall, 1990; meldaHl, 1990; Foote, 

2000; Holland & Patzkowsky, 2002; 

Foote, 2003). Preservation and sampling 

are biased by a number of factors, including 

the rarity and body size of taxa, as well as 

the overall quality and quantity (completeness) 

of the fossil record. The completeness 

of the fossil record varies systematically 

through time with tectonic and/or climatic 

factors (e.g., smitH, Gale, & monks, 2001; 

cramPton, Foote, beu, cooPer, & others, 

2006; S. Peters, 2006). It is also affected by 

the abundance of unlithified versus lithified 

sediments (Hendy, 2009; sessa, Patzkowsky, 

& bralower, 2009). Extinctions that occur 

during intervals of poor preservation and 

sampling appear to happen earlier in time 

(back-smearing), whereas originations in 

those same intervals appear to happen later 

in time (forward-smearing). This problem 

is not unique to studies of the fossil record,

ut rather it presents a general challenge to 

any attempt to estimate extinction (or origination) 

from limited observations (solow, 

1993, 2005; riVadeneira, Hunt, & roy, 

2009). 

The degree of discordance between the 

timing of true extinction and origination 

and the observed stratigraphic ranges of taxa 

depends on temporal variation in preservation 

and sampling. Accounting for such variation 

can be critical in reconstructing diversity 

dynamics over geologic time. Multiple 

approaches have been developed to account 

for variable preservation and sampling 

in paleontological studies. The choice of 

method depends on the specific question 

being addressed and the spatiotemporal 

scale of sampling. For example, at local or 

regional scales, datasets may be partitioned 

to examine only samples collected from 

comparable taphonomic or stratigraphic 

contexts (e.g., scarPoni & kowalewski, 

2007; n. Heim, 2009). At the global scale, 

two general approaches have been taken 

to account for temporal variation in the 

completeness of the known fossil record: 

occurrence-based approaches that rely on 

sub- or replicate-sampling methods, such 

as rarefaction (e.g., alroy & others, 2001; 

busH, markey, & marsHall, 2004; alroy 

& others, 2008) and capture-mark-recapture 

(e.g., connolly & miller, 2002; liow & 

others, 2008), and modeling approaches 

that estimate rates of extinction, origination, 

and preservation simultaneously from the 

observed paleontological data (Foote, 2000, 

2003, 2005). Preservation rate in this last 

approach describes jointly the probability of 

preservation and sampling over time. 

MARINE BIVALVE 

EXTINCTION AND 

ORIGINATION DYNAMICS 

THROUGH THE 

PHANEROZOIC 

Here we estimate extinction, origination, 

and preservation rates simultaneously for 

marine bivalve genera through the Phanero- 


3 

zoic using a likelihood-based modeling 

approach developed by Foote (2003, 2005). 

This approach uses numerical optimization 

to identify the time series of extinction, 

origination, and preservation rates most 

likely to have generated the observed data 

(i.e., the matrices of forward and backward 

survivorship frequencies calculated from 

the observed temporal distribution of first 

and last occurrences of genera) under a 

given model of evolution and preservation. 

Our analysis of bivalve diversity dynamics 

differs from previous studies (e.g., stanley, 

1977; Gould & calloway, 1980; kruG, 

jablonski, & Valentine, 2009), in that 

completeness of the preserved and sampled 

fossil record is explicitly taken into account 

in estimating evolutionary rates. Our analysis 

also focuses on rates of extinction and 

origination rather than diversification rate or 

standing diversity (cf. stanley, 1977; Gould 

& calloway, 1980; miller & sePkoski, 

1988). 

We use a global compilation of observed 

first and last occurrences of marine bivalve 

genera for rate estimation (sePkoski, 2002). 

Data for the 2861 bivalve genera in sePkoski’s 

Compendium of Fossil Marine Animal 

Genera (2002) were compiled primarily from 

the first Treatise on Invertebrate Paleontology 

devoted to the Bivalvia (cox & others, 

1969; stenzel, 1971); data for the revised 

Treatise are as yet unavailable. The Paleobiology 

Database (alroy & others, 2001, 

2008)—a global compilation of spatial and 

temporal occurrences of fossil taxa through 

the Phanerozoic—is another dataset that 

could have been used to investigate bivalve 

evolutionary rates. Although occurrencebased 

data can also be analyzed in such a 

way as to account for variable sampling 

and preservation (see above), we chose to 

analyze sePkoski’s Compendium of first 

and last occurrences to provide a benchmark 

against which analysis of data from 

the revised Treatise could be compared in 

the future. While we are eager to see how 

results differ following the taxonomic revisions 

anticipated in the revised Treatise, we


do not expect substantial changes. Studies 

conducted at comparably broad spatial, 

temporal, and taxonomic scales have shown 

that taxonomic errors tend to be randomly 

distributed and overall macroevolutionary 

patterns are surprisingly robust (adrain & 

westroP, 2000; ausicH & Peters, 2005; 

waGner & others, 2007). 

Rates of extinction, origination, and preservation 

for marine bivalve genera were estimated 

for 71 time intervals that correspond 

roughly to geologic stages. Data for some 

stages were combined to minimize temporal 

variation in interval duration (median interval 

duration = 6.4 million years; interquartile 

range, 4.4 to 10.2 million years). Extinction 

and origination rates were calculated assuming 

a pulsed model of taxonomic turnover in 

which originations cluster at the start of each 

interval and extinctions at the end of each 

interval (Foote, 2003, 2005). Under this 

model, extinction rate equals the number of 

genera last appearing in an interval divided 

by the total diversity of the interval; origination 

rate equals the number of new genera 

in an interval divided by the number present 

at the start of the interval; and preservation 

rate equals the estimated probability of 

being preserved and sampled per genus per 

interval; see Foote (2003) for additional 

methodological details. We have also examined 

extinction and origination rates generated 

under an alternative evolutionary model in 

which taxonomic turnover occurs continuously 

(Foote, 2003). These models—pulsed 

versus continuous turnover—both identify 

peaks in origination and extinction rates for 

marine bivalves through the Phanerozoic, but 

the magnitudes and timing of the peaks differ 

somewhat. We restrict our discussion to the 

results of the pulsed turnover model, as this 

model has greater support globally (Foote, 

2005) and regionally (cramPton, Foote, 

beu, maxwell, & others, 2006), and also 

has the advantage of incorporating all genera, 

including those confined to a single stage, in 

the estimation of all three rates (Foote, 2003). 

Rates of extinction, origination, and preservation 

estimated for marine bivalve genera 

through the Phanerozoic are presented in 

Figure 1. Rates of preservation vary considerably 

over time, spanning nearly the full 

range from zero to one, with a median rate 

equal to 0.33. The median rate of preservation 

for bivalves in this analysis is lower than 

previous estimates (Foote & sePkoski, 1999; 

Valentine & others, 2006). This moderate 

rate of preservation overall, combined with 

its volatility over time, underscores the 

importance of accounting for temporal 

variation in the completeness of the fossil 

record when estimating extinction and 

origination rates. 

In general, bivalves exhibit moderate 

rates of extinction and origination through 

the Phanerozoic (median rate of extinction 

= 0.1; median rate of origination = 0.2), 

with rare intervals of elevated rates (Fig. 

1). Prominent peaks in extinction occurred 

during the late Cambrian, end-Ordovician, 

Late Devonian, end-Permian, end-Triassic, 

and end-Cretaceous, all times of elevated 

extinction observed at much broader taxonomic 

scales (Foote, 2003). These extinction 

peaks have previously been observed 

in studies of the fossil record that assume 

perfect preservation, but it is noteworthy 

that they remain after accounting for the 

dramatic temporal variation observed in 

preservation rate. 

A secular decline in bivalve extinction 

and origination rates over the Phanerozoic 

is observed; the Spearman rank order correlation 

between extinction and origination 

rate and time is 0.26 and 0.25, respectively 

(p-value < 0.05 for both correlation tests). 

The Phanerozoic-scale decline in rates of 

extinction and origination has also been 

observed at broader taxonomic scales (rauP 

& sePkoski, 1982; Van Valen, 1984; Foote, 

2003) and may result from the loss of extinction-prone 

lineages over time (roy, Hunt, 

& jablonski, 2009), although other mechanisms 

have also been proposed (alroy, 2008 

and references therein). Differing taxonomic 

practice could potentially contribute to the 

observed temporal variation in bivalve rates 

of taxonomic extinction and origination.

Extinction rate 

0 0.2 0.4 0.6 0.8 

Origination rate 

0 1.0 2.0 6.0 7.0 

Preservation rate 

0 0.3 0.6 0.9 

However, previous studies conducted at 

comparable scales have generally found 

taxonomic errors to be randomly distributed 

(adrain & westroP, 2000; waGner 

& others, 2007), and there is little reason 


C O S D C P Tr J K Cz 

500 400 300 200 100 0 

Geologic time (Ma) 


500 400 300 200 100 0 



500 400 300 200 100 0 


Extinction rate 

0 0.2 0.4 0.6 0.8 

Origination rate 

0 1.0 2.0 6.0 7.0 

Preservation rate 

0 0.3 0.6 0.9 

0 5 10 15 20 25 30 35 

Frequency 

0 10 20 30 40 50 

Frequency 

0 2 4 6 8 10 12 14 

Frequency 

FiG. 1. Extinction, origination, and preservation rates per interval for marine bivalves through the Phanerozoic. 

The left panel presents the mean time series of rate estimates derived from 100 bootstrap replicate samples of the 

observed data. The right panel presents the frequency distribution of rate magnitudes. Extinction and origination 

rates were estimated assuming a pulsed model of taxonomic turnover. Under this model, extinction rate equals the 

number of genera last appearing in an interval divided by the total diversity of the interval, origination rate equals 

the number of new genera in an interval divided by the number present at the start of an interval, and preservation 

rate is the estimated probability of preservation per genus per interval (new). 

5 

to expect rates of pseudoextinction and 

pseudoorigination to decline toward the 

present day. 

To identify peaks of extinction that stand 

out substantially above the baseline rate for


Extinction rate residuals 

−6 −4 −2 0 2 


500 400 300 200 100 0 


the portion of the Phanerozoic in which 

they occur, the long-term secular trend in 

both evolutionary rates was removed by 

fitting a linear regression to the natural 

logarithm of evolutionary rate against time 

(alroy, 2008). When the residual variation 

in extinction rate is considered, the 

late Cambrian, end-Permian, end-Triassic, 

and end-Cretaceous are times at which 

extinction was particularly severe for marine 

bivalves (Fig. 2). Depending on the cut-off 

value used to define a severe extinction, the 

Eocene–Oligocene and Plio–Pleistocene 

are also noteworthy. This is consistent with 

previous studies that have documented 

substantial molluscan extinction, at least 

regionally, during these times (e.g., raFFi, 

stanley, & marasti, 1985; stanley, 1986a; 

allmon & others, 1993; ProtHero, iVany, 

& nesbitt, 2002; dockery & lozouet, 

2003; Hansen, kelley, & Haasl, 2004). 

Global patterns such as these emerge 

as the sum of processes of extinction and 

origination operating at finer spatial scales. 

Processes of extinction and recovery are 

biogeographically complex. There are 

currently insufficient data to address spatial 

variation in extinction and recovery in any 

detail for many events. Intervals in which 

extinction triggers are both pronounced 

and widespread should result in greater 

congruence between patterns of taxonomic 

Origination rate residuals 


500 400 300 200 100 0 


FiG. 2. Residual variation in extinction and origination rates remaining after removing the secular decline in both 

rates through the Phanerozoic using linear regression. The so-called Big Five mass extinctions recognized in marine 

invertebrates are denoted with large black diamonds (new). 

−6 −4 −2 0 2 

turnover at global and regional scales, but 

such large events are relatively uncommon 

in the history of life. The end-Cretaceous 

(K/Pg, formerly K/T) mass extinction and 

recovery is one example of a large-scale event 

in which the biogeographic fabric of diversity 

loss and rebound has received detailed 

study. At this time, regions differed little 

in the severity of extinction experienced by 

marine bivalves, but markedly in the timing 

and process of recovery (rauP & jablonski, 

1993; jablonski, 1998). Examples of more 

biogeographically differentiated intervals of 

extinction and recovery for marine bivalves 

include the Triassic–Jurassic (e.g., Hallam, 

1981; aberHan, 2002) and Plio–Pleistocene 

(e.g., raFFi, Stanley, & marasti, 1985; 

stanley, 1986a; allmon & others, 1993; 

todd & others, 2002), among others. 

DIVERSITY-DEPENDENT 

DYNAMICS IN THE MARINE 

BIVALVIA THROUGH THE 

PHANEROZOIC 

Over their history, marine bivalves have 

experienced periods of elevated extinction 

and origination, as well as periods 

of relative evolutionary quiescence (Fig. 

1). To what extent have the dynamics of 

extinction and origination been coupled 

over time? Extinction may facilitate origi-

nation through the removal of incumbent 

taxa and opening up of ecospace. Understanding 

whether extinction and origination 

rates operate in a diversity-dependent 

fashion has important implications for our 

understanding of the role of biotic interactions 

in diversification (sePkoski, 1978; 

miller & sePkoski, 1988; kircHner & 

weil, 2000a, 2000b; erwin, 2001; lu, 

yoGo, & marsHall, 2006; alroy, 2008; 

jablonski, 2008a). If diversity-dependent 

dynamics have been important for marine 

bivalves through the Phanerozoic, then 

times of limited extinction should have been 

followed by times of limited origination, 

and times of elevated extinction by times of 

elevated origination. Whether the response 

of origination to extinction was immediate 

or lagged by some period of time depends 

on the nature of the recovery process. If 

extinction empties niches, then origination 

may respond rapidly to new ecological and 

evolutionary opportunity. However, if niches 

depend in part upon diversity and need to 

be reconstructed following major perturbations, 

then temporal lags between extinction 

and origination peaks are to be expected. 

Pseudoextinction—the apparent evolutionary 

turnover of taxa resulting from 

anagenetic morphological evolution and/or 

variation in taxonomic practice—could also 

contribute to a positive relationship between 

extinction and subsequent origination, if 

rates of pseudoextinction are elevated over 

some intervals relative to others. If true, this 

is not necessarily less significant, as pseudoextinction 

presumably reflects some amount 

of morphological change. Thus, temporal 

variation in rates of pseudoextinction could 

offer insight into the evolutionary response 

of taxa to changes in the biotic and abiotic 

environment. In practice, pseudoextinction 

is probably not a major factor governing 

the variation we observe in rates of extinction 

and origination—as well as their association—over 

the Phanerozoic history of 

marine bivalves. Previous studies conducted 

at comparable spatial, temporal, and taxonomic 

scales that have compared taxonomi- 


7 

cally standardized data with data aggregated 

from the literature without taxonomic standardization 

have generally found taxonomic 

errors to be randomly distributed (adrain & 

westroP, 2000; waGner & others, 2007), 

and rate estimates, as a result, to be affected 

little by the process of taxonomic standardization 

(waGner & others, 2007; but see 

ausicH & Peters, 2005). Anagenesis within 

genera cannot be fully accounted for until 

a comprehensive genus-level phylogenetic 

framework exists for the Bivalvia. 

To determine whether evolutionary rates 

were diversity-dependent among marine 

bivalves through the Phanerozoic, we examined 

the variation in rates of extinction 

and origination that remains following the 

removal of the long-term secular decline in 

rates noted above. The effect of extinction 

on origination was evaluated using the slope 

of a linear regression model relating the rate 

of extinction in an interval (t) to the rate 

of origination in the next interval (t + 1). 

We evaluated the support for an effect of 

extinction on subsequent origination by 

assessing whether the observed regression 

slope was significantly greater than zero, 

and whether it differed from that expected 

solely by chance via a permutation test. The 

distribution of null values against which the 

observed slope was compared was generated 

by randomly shuffling the detrended rates of 

extinction and origination and calculating 

the slope of the extinction versus origination 

relationship, and repeating this procedure 

10,000 times. 

A significant positive relationship is 

observed, such that periods of elevated 

extinction are followed by elevated origination, 

and periods of moderate extinction 

are followed by moderate origination (Fig. 

3; Table 1). The results of the permutation 

test also indicate that the observed relationship 

between extinction rate and subsequent 

origination rate among marine bivalves is 

significantly greater than expected by chance 

(Fig. 4). There is no indication of a lag in 

the response of origination to extinction; 

rather, origination responds immediately in


Origination rate (t+1) 

−6 −4 −2 0 2 

−6 −4 −2 

Extinction rate (t) 

0 2 

FiG. 3. The effect of extinction on origination for marine 

bivalve genera through the Phanerozoic. Plotted 

are the rates of extinction in each interval (t) against 

origination in the next interval (t + 1). Rates were logarithmically 

transformed and detrended prior to analysis; 

details provided in text. The dashed line is a linear 

regression model fitted to the extinction-origination 

relationship. A statistically significant positive relationship 

is observed, indicating that diversity-dependent 

processes have operated over the evolutionary history 

of the Bivalvia (new). 

the next interval, and this effect subsequently 

weakens over time. The association between 

extinction rate in an interval (t) and origination 

rate in the next interval (t + 1) was 

approximately double that of extinction rate 

and origination rate two intervals later (t + 2) 

(i.e., slopes of 0.30 and 0.16 respectively). 

These results are consistent with studies of 

the relationship between extinction and origination 

for skeletonized marine invertebrates 

as a whole (lu, yoGo, & marsHall, 2006; 

alroy, 2008), and corroborate previous 

work on marine bivalves that documented 

hyperexponential bursts of diversification 

following mass extinction events (miller & 

sePkoski, 1988; kruG, jablonski, & Valentine, 

2009). It is important to note, however, 

that the diversity-dependent relationship 

between rates of extinction and origination 

for marine bivalves is not limited to mass 

extinctions and their associated recoveries. 

While removing the most extreme extinction 

events from the analysis somewhat 

weakens the relationship between extinction 

and subsequent origination, intervals char- 

table 1. Effect of extinction on origination for 

marine bivalve genera through the Phanerozoic. 

Effect was measured as the slope of the 

linear regression of extinction rate in an interval 

(t) on origination rate in the next interval 

(t + 1). Rates were detrended prior to analysis; 

details provided in text. A significant positive 

relationship is observed; intervals of elevated 

extinction are followed by intervals of elevated 

origination, and intervals of moderate extinction 

followed by intervals of moderate origination. 

This diversity-dependent relationship between 

extinction and origination is not driven 

simply by mass extinctions and their associated 

recovery intervals. Excluding intervals characterized 

by elevated extinction (e.g., the top 

5% of extinctions, top 10%) does not markedly 

weaken the overall relationship (new). 

Data Effect p-value 

All 0.3 0.02 

excluding top 5% 0.25 0.07 




acterized by relatively low extinction rates 

also exhibit diversity dependence (Table 1). 

Extinction has been an important evolutionary 

process throughout the history of 

marine bivalves, varying considerably in 

intensity over time, but contributing consistently 

to bivalve diversification, in part, 

through its effect on rates of origination. 

INFLUENCE OF BIOLOGICAL 

FACTORS ON EXTINCTION 

RISK AMONG MARINE 

BIVALVES 

Extinction selectivity, or the selective 

removal of taxa that possess particular 

ecological or evolutionary traits, also plays 

an important role in shaping macroevolutionary 

and macroecological patterns 

through time. Extinction selectivity can 

contribute to ecosystem reorganization by 

eliminating dominant taxa and allowing 

subordinate ones to diversify (Gould & 

calloway, 1980; jablonski, 1986, 1989; 

droser, bottjer, & sHeeHan, 1997); it can

edirect evolutionary or ecological trends by 

eliminating important innovations (Pojeta 

& Palmer, 1976; FürsicH & jablonski, 

1984; jablonski, 1986); and it can limit the 

potential evolution of clades by reducing 

variability (norris, 1991; liu & olsson, 

1992). By studying extinction selectivity 

over long time scales, one can assess not 

only which taxa went extinct, but potentially 

how and why they did so. This link 

between extinction pattern and process can 

help to bridge the gap between paleontology 

and conservation biology (see papers in 

dietl & Flessa, 2009). If we can determine 

which traits have influenced susceptibility to 

extinction during periods of past environmental 

change, then we may be better able 

to predict which organisms are most likely 

to go extinct or persist in the present day. 

Extinction selectivity is thought to have 

significantly influenced the evolutionary 

trajectory of marine invertebrates and the 

ecological structure of marine ecosystems 

through geologic time. Bivalves are no 

exception; indeed, several classic studies 

of extinction selectivity have focused on 

the long and relatively well-preserved fossil 

record of marine bivalves (e.g., bretsky, 

1973; kauFFman, 1978; jablonski, 1986; 

stanley, 1986a; jablonski, 2005). This 

is in part because bivalves display sufficient 

variation among taxa in traits such as 

morphology, feeding mode, life habit, larval 

type, geographic range, and stratigraphic 

range to allow workers to independently 

test the extent to which these traits relate to 

taxon survivorship. 

A review of the literature on extinction 

selectivity in fossil marine bivalves published 

in 2010 and before (see Appendix, p. 20) 

demonstrates that selectivity among taxa 

has been explored with respect to a wide 

variety of traits, such as abundance, feeding 

mode, life habit, geographic range, body 

size, temperature tolerance, species richness, 

and habitat breadth, among many others 

(33 traits in total). This review includes 170 

tests of extinction selectivity published in 

69 studies. The vast majority of selectivity 

studies have focused on Mesozoic (120 


Frequency 

0 500 1000 1500 

−0.6 −0.3 0.0 0.3 0.6 

Effect of extinction (t) on origination (t+1) 

9 

FiG. 4. The effect of extinction on subsequent origination 

for marine bivalve genera through the Phanerozoic. 

Effect was measured using the slope of the linear regression 

of extinction rate in an interval (t) on origination 

rate in the next interval (t + 1). The gray frequency 

distribution presents the randomized values, the solid 

arrow denotes the observed effect, and the dashed line 

indicates the 95th quantile of randomized values. The 

observed effect is significantly greater than expected by 

chance (new). 

tests) and Cenozoic (114 tests) bivalves, 

with the Paleozoic receiving considerably 

less attention (18 tests). The extinctions 

represented in our database range from the 

largest and most catastrophic mass extinctions 

(including all of the so-called Big Five), 

to six smaller, possibly regional-scale, events 

(e.g., Eocene/Oligocene and Plio/Pleistocene) 

and background intervals. Selectivity 

has been examined at both the species (98 

tests) and genus (80 tests) levels. Examining 

the specific traits tested for selectivity, four 

traits have received the most attention: 

geographic range (27 tests), life habit (28 

tests), body size (21 tests), and feeding mode 

(15 tests). 

If extinction is defined as the point in 

time at which a taxon’s geographic range 

and abundance decrease to zero, taxa 

with broader geographic ranges should 

be less prone to extinction. The primary 

role that geographic distribution plays in 

determining survivorship has long been 

recognized for both modern and fossil taxa 

(see references in Gaston, 1994; rosenzweiG, 

1995; Payne & FinneGan, 2007). 

An early assessment of global survivorship


across four mass extinctions—the end- 

Ordovician, Late Devonian, end-Permian, 

and end-Triassic events—concluded that 

geographically widespread bivalve genera 

were more likely to survive, at least in the 

initial stages of an extinction event, before 

drastic deterioration of the physical environment 

(bretsky, 1973). The event that 

has been most thoroughly examined for 

geographic range selectivity is the K/Pg mass 

extinction, in conjunction with the interval 

of background extinction leading up to it. 

K/Pg survivorship patterns in bivalves and 

gastropods, along the United States Gulf 

and Atlantic Coastal Plain and globally, 

suggest that the only trait reliably associated 

with genus survivorship is geographic 

range, whereas several traits are associated 

with genus- and species-level survivorship 

during the preceding background interval 

(jablonski, 1986, 1987, 1989; rauP & 

jablonski, 1993; jablonski & rauP, 1995; 

jablonski, 2005; jablonski & Hunt, 2006). 

Data for Southern Hemisphere bivalves 

across the K/Pg, also compiled at the genus 

level, support this general pattern (stilwell, 

2003). In contrast, during the end-Triassic 

extinction, European bivalve species with 

broader distributions appear no more likely 

to have survived this event than narrowly 

distributed taxa (mcroberts & newton, 

1995; mcroberts, newton, & allasinaz, 

1995). When the spatial scale of 

analysis is expanded to global coverage, the 

same nonsignificant species-level pattern is 

observed for the end-Triassic (kiesslinG & 

aberHan, 2007). 

Taxonomic level and the intensity 

of extinction complicate the pattern of 

geographic range selectivity. The data 

compiled here strongly suggest that widespread 

bivalve species were significantly more 

likely to survive background (jablonski, 

1986, 1987; jablonski & Hunt, 2006; 

kiesslinG & aberHan, 2007; cramPton 

& others, 2010; Harnik, 2011), but not 

mass extinction events (jablonski, 1986; 

Hansen & others, 1993; mcroberts & 

newton, 1995; mcroberts, newton, & 

allasinaz, 1995; jablonski, 2005; kiesslinG 

& aberHan, 2007; for exceptions, see rode 

& lieberman, 2004, for the Late Devonian, 

and stilwell, 2003, for the K/Pg). Late 

Neogene extinctions, intermediate in scale 

between the Big Five events and background 

extinction, differ in the effects of geographic 

range on selectivity among regions. 

Narrowly distributed species were more 

likely to become extinct in western South 

America (riVadeneira & marquet, 2007) 

and tropical America (rooPnarine, 1997), 

but not in western North America, where 

there is no evidence of selective extinction 

(stanley, 1986b). These complex patterns 

highlight the importance of assessing selectivity 

across a range of extinctions that differ 

in magnitude, as well as across a range of 

taxonomic levels. It is possible that thresholds 

exist, such that geographic range no 

longer ensures the survival of a species, when 

the scale of environmental perturbation 

and extinction exceed a critical magnitude. 

If true, this has enormous implications for 

assessment of extinction risk and development 

of effective management strategies in 

modern ecosystems. 

Bivalve life habit, specifically the sites the 

animals occupy relative to the sedimentwater 

interface, is another ecological trait 

that is thought to affect survivorship. 

Which life habit favors survival probably 

depends on the mechanism of extinction. 

For example, epifaunal taxa are thought to 

be more vulnerable to predation pressure 

(stanley, 1977, 1982, 1986b; Vermeij, 

1987), which could lead to decreased population 

size and an increase in extinction risk. 

In contrast, an epifaunal life habit may be 

advantageous in escaping sudden changes in 

bottom water chemistry and/or oxygenation. 

Of the studies compiled here, 14 suggested 

that epifaunal taxa were more likely to 

become extinct than infaunal, 5 suggested 

the opposite, and 8 found no evidence of 

selectivity either way. When these results are 

parsed according to the size of the extinction 

event, an interesting pattern emerges. The 

majority of background intervals studied 

(8 out of 10) suggest that infaunal bivalves 

were less likely to go extinct, while mass

extinctions yield contradictory results (4 

negative, 5 positive, 6 nonsignificant). When 

the mass extinction events are broken down 

by specific event, the results remain mixed. 

For example, while bivalve genera globally 

did not exhibit differential survival 

with respect to life habit across the end- 

Permian mass extinction (jablonski, 2005), 

regional patterns from China suggested 

greater losses of epifaunal than infaunal 

bivalve genera (knoll & others, 2007). 

Perhaps these differences reflect the extent 

to which different geographic regions were 

affected by environmental deterioration. In 

another example, preferential extinction of 

infaunal bivalve species was documented 

across the K/Pg boundary in New Jersey 

and the Delmarva Peninsula of the United 

States (GallaGHer, 1991), but subsequent 

work found the opposite pattern for bivalve 

species in Denmark (HeinberG, 1999) and 

the Southern Hemisphere (stilwell, 2003). 

Although global analyses of selectivity 

can be very useful in seeking possible causes 

of extinction, they can obscure regional 

patterns that may be less predictable and yet 

likely to provide more information about 

the interacting effects of biotic and abiotic 

factors on survivorship. Spatial variation in 

environmental change, coupled with spatial 

heterogeneity in the distributions of taxa and 

associated biological traits, effectively ensure 

that patterns of selectivity will vary regionally 

(see Fritz, bininda-emonds, & PurVis, 

2009, for an example of geographic variation 

in extinction risk among extant mammals). 

Spatial variation may provide useful information 

about gradients of environmental change 

and the existence of environmental thresholds 

affecting taxon survivorship. Despite the 

clear importance of regional-scale studies in 

modern conservation biology, paleontological 

examples are few and far between. 

Although large body size is widely thought 

to increase extinction risk in vertebrates, 

the link between size and extinction risk in 

marine invertebrates is considerably more 

ambiguous (Hallam, 1975; stanley, 1986b; 

budd & joHnson, 1991; jablonski, 1996b; 

smitH & roy, 2006). Among invertebrates, 


11 

increased body size is often associated 

with increased fecundity, broader environmental 

tolerance, and wider geographic 

range (stanley, 1986b; mckinney, 1990; 

rosenzweiG, 1995; Hildrew, raFFaelli, & 

edmonds-brown, 2007), which suggests 

that larger taxa should have increased rates 

of survivorship. Among marine bivalves, 

however, large body size is not generally associated 

with either extinction risk or survivorship. 

Body size and extinction are positively 

linked in only 7 and negatively linked in only 

2 (out of a total of 21) studies. Four of the 7 

studies that documented selective extinction 

of large taxa focused on regional patterns 

during intervals characterized by background 

rates of extinction; these include the Jurassic 

(Hallam, 1975), Miocene (anderson & 

rooPnarine, 2003, for the Western Atlantic 

and Caribbean, but not the Eastern Pacific), 

and Pleistocene (stanley, 1986a, 1990b). 

Interestingly, not a single one of the 10 

studies that considered size across a mass 

extinction event found a strong, conclusive 

link between body size and extinction, 

although the only 2 events investigated 

thus far are the end-Triassic (mcroberts & 

newton, 1995; mcroberts, newton, & 

allasinaz, 1995; mcroberts, krystyn, & 

sHea, 2008) and K/Pg (Hansen & others, 

1987; rauP & jablonski, 1993; jablonski 

& rauP, 1995; mcclure & boHonak, 

1995; jablonski, 1996a; lockwood, 2005; 

aberHan & others, 2007) events. Three 

of these 10 studies documented a decrease 

in bivalve size across a mass extinction 

boundary (Norian–Rhaetian: mcroberts, 

krystyn, & sHea, 2008; K/Pg: Hansen & 

others, 1987; aberHan & others, 2007), 

but it is unclear in each case whether these 

patterns were driven by extinction selectivity, 

within lineage size change, or size-biased 

origination. 

One of the few instances in which a 

connection between large body size and 

survivorship has been documented convincingly 

focused on scallops across the Plio– 

Pleistocene extinction in California (smitH 

& roy, 2006). This positive relationship was 

not apparent until phylogenetic relationships


were considered. This emphasizes an underappreciated 

problem that may affect many 

selectivity studies. Patterns of selectivity can 

sometimes be masked or artificially exaggerated 

when phylogenetic relationships are not 

taken into account (PurVis, 2008). Taxa may 

share a particular trait and similar pattern 

of survivorship because they are related 

to each other and not necessarily because 

the trait under consideration, by itself, 

confers survivorship. A recent analysis (roy, 

Hunt, & jablonski, 2009) of Jurassic to 

Recent bivalves demonstrated conclusively 

that phylogenetic clustering of extinction 

occurs. Phylogenetic relationships do not 

always affect patterns of selectivity, however; 

for example, patterns of selectivity among 

Cenozoic mollusks from New Zealand did 

not change appreciably after accounting 

for phylogeny (Foote & others, 2008; 

cramPton & others, 2010). The fact that 

taxa in some clades are significantly more 

extinction-prone than others does strongly 

suggest that future paleontological studies 

of selectivity should explicitly account for 

phylogenetic effects. 

In general, deposit feeding is thought 

to represent a more generalized dietary 

mode than suspension feeding and could 

promote survivorship, especially across 

events that involve a collapse in primary 

productivity. leVinton’s (1974) observation 

that genera of deposit-feeding bivalves were 

geologically longer-lived than suspensionfeeders, 

has inspired an ongoing debate 

over whether bivalves with different feeding 

modes experience different extinction trajectories. 

Building on this work, a qualitative 

examination of background extinction in 

Cretaceous bivalve species documented 

particularly slow rates of evolution and long 

stratigraphic durations in deposit-feeders 

relative to suspension-feeders (kauFFman, 

1978). The majority of studies that have 

explicitly tested for extinction selectivity 

according to feeding mode have focused on 

the K/Pg event, with mixed results. Seven 

out of 11 studies have reported selective 

extinction of suspension-feeding genera 

(e.g., sHeeHan & Hansen, 1986; rHodes 

& tHayer, 1991; rauP & jablonski, 1993; 

jablonski & rauP, 1995; jablonski, 1996a; 

stilwell, 2003; aberHan & others, 2007). 

There is preliminary evidence to suggest 

that the strength of the selectivity may 

have increased with distance away from the 

United States Gulf Coastal Plain, which 

raises the question as to whether proximity 

to the killing agent, in this case the K/Pg 

bolide impact that occurred in the Yucatan, 

has an effect on selectivity patterns. Studies 

limited to eastern Texas (Hansen & others, 

1987; Hansen, Farrell, & uPsHaw, 1993; 

Hansen & others, 1993, for exception see 

sHeeHan & Hansen, 1986) or the United 

States Gulf and Atlantic Coastal Plain 

(mcclure & boHonak, 1995), have yielded 

either weak or no evidence for selectivity. 

On the other hand, regional studies in the 

Southern Hemisphere (e.g., stilwell, 2003) 

and Argentina (aberHan & others, 2007) 

have suggested that proximity matters, as 

they have shown strong evidence for selectivity. 

Although jablonski’s work (rauP & 

jablonski, 1993; jablonski & rauP, 1995; 

jablonski, 1996a) clearly supports a global 

pattern of selective extinction of suspensionfeeding 

bivalves at the K/Pg, he has argued 

that this was driven by taxonomic factors, 

rather than selectivity according to feeding 

mode. He and his colleagues pointed to 

anomalously low rates of extinction in the 

two bivalve orders Nuculoida and Lucinoida 

and argued that other attributes of these two 

clades helped to promote their survivorship. 

sHeeHan and Hansen (1986), Hansen and 

others (1987, 1993), and Hansen, Farrell, 

and uPsHaw (1993) emphasized the shift 

from molluscan communities dominated by 

suspension-feeders to those dominated by 

deposit-feeders across the K/Pg boundary, a 

pattern that could have been caused by selective 

extinction against suspension-feeders, 

preferential recovery of deposit-feeders, 

or some combination of the two. Explicit 

evaluation of this possibility, in addition to 

detailed tracking of feeding mode across the 

recovery interval, is still lacking. Although 

early studies heralded the usefulness of 

patterns of extinction selectivity based on

feeding habits in differentiating among 

possible extinction mechanisms, this potential 

has seldom been realized (but see knoll 

& others, 1996, 2007, for exceptions). As 

our understanding of changes in primary 

productivity associated with mass extinctions 

deepens, aided by geochemical proxies, 

it should be possible to further refine and 

test hypotheses bearing on the relationship 

between feeding mode and extinction risk 

across an array of marine environments. 

Most of the studies outlined above focus 

on the selectivity of single traits and do not 

consider the potential interactions among 

multiple traits. We have every reason to 

believe, based on ecological studies of extant 

bivalves and many other clades, that several 

of these traits, for example, body size and 

geographic range (jablonski & roy, 2003; 

cramPton & others, 2010; Harnik, 2011), 

are linked to one another. This raises the 

question—to what extent do these interactions 

influence patterns of selectivity? A 

handful of recent studies have tackled this 

question for marine bivalves (jablonski & 

Hunt, 2006; riVadeneira & marquet, 

2007; jablonski, 2008a; cramPton & 

others, 2010; Harnik, 2011). Almost all 

of them have found that geographic range 

played a more important role in survivorship 

than any other ecological trait. For example, 

in a genus-level analysis of selectivity across 

the K/Pg mass extinction, jablonski (2008a) 

independently tested the effects of body 

size, geographic range, and species richness, 

and found that the last two traits were both 

statistically significantly correlated with 

survivorship. However, once the covariation 

among these three traits was controlled for, 

geographic range yielded the only significant 

evidence for selectivity. In what is perhaps 

the most extensive multivariate selectivity 

study to date, cramPton and others (2010) 

assessed the relative importance of several 

traits, including geographic range, body size, 

feeding mode, life habit, and larval type, in 

promoting survivorship among Cenozoic 

bivalve species from New Zealand. Once 

again, in a multivariate framework, the 

only trait to show demonstrable selectivity 


13 

was geographic range. Such multivariate 

approaches are crucial to studies of selectivity, 

offering considerable insight into the 

direct and indirect effects of extinction on 

the evolution of correlated traits. A clear 

understanding as to how traits interact, 

influencing extinction risk across a range of 

past events, is needed, if bivalve workers are 

to make such patterns relevant to managers 

predicting biotic response to current extinction 

pressures. 

CONCLUSIONS 

One of the major insights of paleontology 

is the importance of extinction in shaping 

the diversity of life through time. The 

effects of extinction on diversity dynamics 

have been intensively studied in the marine 

Bivalvia because of their relatively complete 

fossil record, the considerable biological 

variation that exists among taxa, and their 

diversity and abundance in shallow marine 

environments today and in the past. In this 

contribution, we provide new estimates of 

global extinction and origination rates for 

marine bivalve genera through the Phanerozoic 

that explicitly account for temporal 

variation in preservation. These analyses, 

using data compiled primarily from the first 

Treatise on Invertebrate Paleontology (Part 

N: cox & otHers, 1969; stenzel, 1971), 

underscore the important contributions of 

the Treatise to our understanding of bivalve 

macroevolutionary history. While rates of 

extinction and origination are moderate 

for marine bivalves overall, times of severe 

extinction and times of general quiescence 

are observed through the Phanerozoic. Intervals 

of elevated global extinction for marine 

bivalves correspond to intervals of elevated 

extinction for marine invertebrates more 

broadly, and bivalves exhibit secular declines 

in rates of extinction and origination over 

the Phanerozoic that are also observed at 

much broader taxonomic scales. Throughout 

their history, marine bivalves exhibited 

coupled dynamics of extinction and origination, 

with periods of elevated extinction 

followed by periods of elevated origination, 

and moderate extinction by moderate origi-


nation. This diversity-dependent process is 

most pronounced following mass extinctions, 

but operated consistently throughout 

the history of the clade. Studies of marine 

bivalves have yielded important insights into 

extinction selectivity, and specifically, the 

effects of biological traits on survivorship. 

We review this literature, focusing on four 

traits that have received the most attention. 

Geographic range size is the most consistent 

predictor of bivalve survivorship considered 

to date. Traits like feeding mode and life 

habit may also be important, but these are 

probably more dependent on the particular 

context of environmental change. Body 

size is largely decoupled from extinction 

risk despite reasons to expect otherwise. 

The growing paleontological literature on 

selectivity underscores the major contribution 

of fossil bivalves to our understanding 

of the factors that influence extinction risk. 

It highlights a fruitful area for collaboration 

between researchers studying the effects of 

extinction on marine systems today and in 

the past. 

ACKNOWLEDGMENTS 

Michael Foote generously provided the 

rate estimates presented in Figure 1, and 

his assistance throughout the preparation 

of this manuscript was invaluable. 

Thank you to Heather Richardson at the 

College of William and Mary for her help 

in compiling the bibliography. The quality 

of this manuscript was greatly improved by 

the comments of Martin Aberhan, Joseph 

Carter, James Crampton, Patricia Kelley, 

Roger Thomas, and members of Jonathan 

Payne’s lab at Stanford University. A portion 

of this work was conducted while Rowan 

Lockwood was a Sabbatical Fellow at the 

National Center for Ecological Analysis and 

Synthesis, a Center funded by NSF (Grant 

#EF-0553768), the University of California, 

Santa Barbara, and the State of California. 

This material is based upon work supported 

by the National Science Foundation under 

Grant No. EAR-0718745 to Rowan Lockwood 

and collaborators. 

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Canada. Fisheries 18(9):6–22.


aPPendix. Database of extinction selectivity studies in fossil marine bivalves; please note that the compilation of past work summarized here is ongoing 

and should not be considered exhaustive (new). 

Trait Reference Pattern Taxon Taxon Age Location Notes 

level 

Abundance Harnik, 2011 NS 3 superfm sp Eo US Gulf CP 

Abundance Kiessling & Aberhan, 2007 NS biv sp T/J Global 

Abundance Kiessling & Aberhan, 2007 Neg biv sp Tri-Jur Global Geo range more important than abundance in determining extinction risk. 

Abundance Lockwood, 2003 NS biv gen K/Pg US CP 

Abundance McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Abundance Simpson & Harnik, 2009 Nonlinear biv gen post Pz Global Rare and abundant genera exhibit elevated extinction rates. 

Active locom Aberhan & others, 2007 Pos? moll dom sp K/Pg Argentina Decline of individuals with active lifestyles across boundary, excluding 

nuculoids and lucinids. May or may not be due to extinction selectivity. 

Active locom Rhodes & Thayer, 1991 Pos biv gen K/Pg Global 

Aragonite Hautmann & others, 2008 Pos biv dom sp T/J Tibet 

Bathy range Jablonski & Raup, 1995 NS biv gen K/Pg US CP 

Bathy range Jablonski, 1996a NS biv, gast gen K/Pg Global 

Bathy range Raup & Jablonski, 1993 NS biv gen K/Pg Global 

Body size Aberhan & others, 2007 Pos? moll dom sp K/Pg Argentina Size decrease may or may not be due to extinction selectivity. 

Body size Anderson & Roopnarine, 2003 Pos corbulids gen Mio Carib/W Atl 

Body size Anderson & Roopnarine, 2003 NS corbulids gen Mio E Pacific 

Body size Crampton & others, 2010 NS biv sp Cz NZ 

Body size Hallam, 1975 Pos biv, ammon gen, sp Jur W Euro 

Body size Hansen & others, 1987 Pos? biv sp K/Pg E Texas Shift to smaller bivalves. May or may not be extinction selectivity. 

Body size Harnik, 2011 Varies 3 superfm sp Eo US Gulf CP Correlation between size and duration positive in veneroids, negative in 

pectinoids, not significant in carditoids 

Body size Jablonski & Raup, 1995 NS biv gen K/Pg Global 

Body size Jablonski, 1996a NS biv gen K/Pg Global 

Body size Jablonski, 1996b NS biv, gast sp Late Cret US CP 

Body size Lockwood, 2005 NS 3 groups gen E/O N Am, Euro Large taxa radiate after extinction. Groups include veneroids, arcticoids, 

and glossoids. 

Body size Lockwood, 2005 NS 3 groups gen K/Pg N Am, Euro Small taxa radiate after extinction. Groups include veneroids, arcticoids, 

and glossoids. 

Body size McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Body size McRoberts & Newton, 1995 NS biv sp T/J Euro 

Body size McRoberts, Newton, & Allasinaz, 1995 NS biv sp T/J Euro 

Body size McRoberts, Krystyn, & Shea, 2008 Pos? Monotis sp Nor/Rhaet Austria Size decrease may or may not be due to extinction selectivity. 

Body size Raup & Jablonski, 1993 NS biv gen K/Pg Global 

Body size Smith & Roy, 2006 Neg scallops gen Plio/Pleist California Pattern significant only when adjusted for phylogenetic bias. 

Body size Stanley, 1986a Neg biv sp Plio/Pleist SE US Siphonate bivalves only. 

Body size Stanley, 1986a Pos biv sp Pleist-Rec W N Am, Japan Siphonate bivalves only. 

Body size Stanley, 1990b Pos biv sp Pleist-Rec W N Am, Japan 

Bur depth Kauffman, 1978 Neg biv sp Cret N Amer Suspension feeders only. 

Bur depth Lockwood, 2004 NS 3 groups gen K/Pg N Am, Euro Deeper burrowing taxa radiate during recovery.

aPPendix (Continued). 


Bur depth McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Bur depth Stanley, 1990b Neg biv sp Pleist-Rec W N Am, Japan 

Endemicity Aberhan & Fürsich, 1997 Pos biv sp Pliens-Toar Andean basin 

Endemicity Aberhan & Fürsich, 2000 Pos biv sp Pliens-Toar Andean basin 

Endemicity Hallam, 1981 Neg biv gen T/J Global 

Endemicity Stanley, 1986a Pos biv sp Plio/Pleist SE US 

Endemicity Vermeij, 1986 Pos moll sp Plio N Atl 

Environment Rode & Lieberman, 2004 See notes biv, brach sp F/F N Amer Increased survival for taxa inhabiting middle-outer platform environments, 

but not taxa inhabiting open shelf. 

Epifaunal Aberhan & Baumiller, 2003 Neg biv sp early Jur NW Euro, Andes 

Epifaunal Aberhan & Fürsich, 1997 Pos? biv sp Pliens-Toar Andean basin Long-term increase in infaunal suspension feeders may be due to extinction 

selectivity or environmental change. Both infaunal and epifaunal 

suspension-feeders decrease across boundary. 

Epifaunal Clapham & Bottjer, 2007 Pos? biv, gast gen Perm Global Increase in abundance of infaunal bivalves may or may not be due to 

extinction selectivity. 

Epifaunal Cope, 2002 Pos biv gen O/S Global 

Epifaunal Crame, 2002 Pos biv gen, sp Cret Global In higher latitudes. 

Epifaunal Crampton & others, 2010 NS biv sp Cz NZ 

Epifaunal Gallagher, 1991 Neg moll sp K/Pg US Atl CP 

Epifaunal Hautmann & others, 2008 Neg biv dom sp T/J Tibet 

Epifaunal Heinberg, 1999 Pos? biv sp K/Pg Denmark Within habitat comparison shows increase in infaunal species across 

boundary, may or may not be due to extinction selectivity. 

Epifaunal Jablonski & Raup, 1995 NS biv gen K/Pg Global Excluding rudists. 

Epifaunal Jablonski, 1996a NS biv gen K/Pg Global Excluding rudists. 

Epifaunal Jablonski, 2005 NS biv gen K/Pg Global 

Epifaunal Jablonski, 2005 NS biv gen P/T Global 

Epifaunal Jablonski, 2005 Pos biv gen T/J-K/Pg Global 

Epifaunal Kauffman, 1977 Pos biv sp Cret W Interior 

Epifaunal Kauffman, 1978 Pos biv sp Cret N Amer 

Epifaunal Knoll & others, 2007 Pos biv gen P/T China 

Epifaunal Levinton, 1973 Pos 6 species sp Rec New York Slower evolutionary rates among infaunal and deep water epifaunal, relative 

to shallow water epifaunal taxa. 

Epifaunal Levinton, 1974 Pos 4 groups gen Phan Global Comparison of mortality rates between selective suspension-feeders 

(Pectinacea, Pteriacea, Veneracea) and detritus feeders (Nuculoida). 

Epifaunal Mander & Twitchett, 2008 - biv gen T/J NW Euro Poor preservation of early Jurassic infaunal taxa indicates that selectivity 

may be artifactual. 

Epifaunal McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Epifaunal McRoberts & Newton, 1995 Neg biv sp T/J Euro 

Epifaunal McRoberts, 2001 Pos biv gen T/J Global 

Epifaunal McRoberts, Newton, & Allasinaz, 1995 Neg biv sp T/J Euro 

Epifaunal Raup & Jablonski, 1993 NS biv gen K/Pg Global 

Epifaunal Rivadeneira & Marquet, 2007 Pos biv sp Late Ng W S Amer 

21




level 

Epifaunal Stanley, 1986a NS biv sp Plio/Pleist SE US 

Epifaunal Stilwell, 2003 Pos moll gen, sp K/Pg S hemis 

Escalation Hansen & others, 1999 NS biv, gast sp E/O US CP Used morphology as proxy for escalation. 

Escalation Hansen & others, 1999 NS biv, gast sp K/Pg US CP Used morphology as proxy for escalation. 

Escalation Hansen & others, 1999 NS biv, gast sp Mio US CP Used morphology as proxy for escalation. 

Escalation Hansen & others, 1999 Pos biv, gast sp Plio/Pleist US CP Used morphology as proxy for escalation. 

Escalation Reinhold & Kelley, 2005 NS biv, gast gen, sp K/Pg Mississippi Used morphology as proxy for escalation, only one trait (crenulate 

margins) associated with higher extinction risk in bivalves. 

Geo range Bretsky, 1973 Neg biv gen F/F Global 

Geo range Bretsky, 1973 Neg biv gen O/S Global 

Geo range Bretsky, 1973 Neg biv gen P/T Global 

Geo range Bretsky, 1973 Neg biv gen T/J Global 

Geo range Crampton & others, 2010 Neg biv sp Cz NZ 

Geo range Hallam & Wignall, 1997 Neg biv gen T/J Global 

Geo range Hansen & others, 1993 NS moll sp K/Pg E Texas 

Geo range Harnik, 2011 Neg 3 superfm sp Eo US Gulf CP 

Geo range Jablonski & Hunt, 2006 Neg moll sp Cret-Cz US CP Geo range is more important than larval mode in predicting extinction risk. 

Geo range Jablonski & Raup, 1995 Neg biv gen K/Pg Global 

Geo range Jablonski, 1986 NS biv, gast gen K/Pg US CP 

Geo range Jablonski, 1986 NS biv, gast sp w/in gen K/Pg US CP 

Geo range Jablonski, 1986 Neg biv, gast sp w/in gen Late Cret US CP 

Geo range Jablonski, 1987 Neg biv, gast sp Cret US CP 

Geo range Jablonski, 1989 Neg biv, gast gen K/Pg N Am 

Geo range Jablonski, 2005 NS biv, gast sp w/in gen K/Pg US CP 

Geo range Jablonski, 2005 Neg biv, gast sp w/in gen Late Cret US CP Genera containing widespread species more extinction resistant. 

Geo range Kiessling & Aberhan, 2007 NS biv sp T/J Global 

Geo range Kiessling & Aberhan, 2007 Neg biv sp Tri-Jur Global Geo range more important than abundance in determining extinction risk. 

Geo range McRoberts & Newton, 1995 NS biv sp T/J Euro 

Geo range McRoberts, Newton, & Allasinaz, 1995 NS biv sp T/J Euro 

Geo range Raup & Jablonski, 1993 Neg biv gen K/Pg Global 

Geo range Rivadeneira & Marquet, 2007 Neg biv sp Late Ng W S Amer 

Geo range Rode & Lieberman, 2004 Neg biv, brach sp F/F N Amer 

Geo range Roopnarine, 1997 Neg chionines sp Late Ng trop Amer Regions with higher degrees of endemism affected more severely. 

Geo range Stanley, 1986b NS biv sp Late Ng W N Amer 

Geo range Stilwell, 2003 Neg moll gen, sp K/Pg S hemis 

Hypercapnia Bottjer & others, 2008 Pos? biv gen P/T Global 4 cosmopolitan bivalve genera flourish after P/T, probably tolerant of 

H S and CO , may or may not be due to extinction selectivity. 

2 2 

Hypercapnia Knoll & others, 1996 Pos biv gen P/T China 

Hypercapnia Knoll & others, 1996 Pos biv gen P/T China 

Invaders Rode & Lieberman, 2004 Neg biv, brach sp F/F N Amer



Latitude Crame, 2002 Pos biv gen, sp Cret Global Epifaunal taxa only. 

Meta rate Knoll & others, 2007 Neg biv gen P/T China 

Morph com Kauffman, 1978 NS biv sp Cret N Amer 

Morph var Kolbe, Lockwood, & Hunt, 2011 Neg veneroids sp Late Ng Florida 

Multivariate Crampton & others, 2010 See notes biv sp Cz NZ Geographic range significant. Epifaunal, suspension-feeder, body size, and 

planktonic larva not significant. 

Multivariate Harnik, 2011 See notes 3 superfm sp Eo US Gulf CP Geographic range strong. Abundance weak. Body size weak overall but 

can be as strong as geographic range in specific superfamilies. 

Multivariate Jablonski & Hunt, 2006 See notes moll sp Cret-Cz US CP Geographic range more important than larval type. 

Multivariate Jablonski, 2008a See notes biv gen K/Pg Global Geographic range significant. Body size and species richness nonsignificant. 

Multivariate Rivadeneira & Marquet, 2007 See notes biv sp Late Ng W S Amer Epifaunal, geographic range, body size significant. 

Occupancy Foote & others, 2007 Neg moll sp Cz NZ Species at greatest risk are those that have already been in decline for a 

substantial period of time. 

Pachyodont Jablonski, 2008a Pos rudists gen K/Pg Global May be example of trait hitch-hiking. Geographic range is more important 

in determining risk. 

Phylogenetic Roy, Hunt, & Jablonski, 2009 Pos biv fam, gen Jur-Rec Global 

Plank larva Crampton & others, 2010 NS biv sp Cz NZ 

Plank larva Gallagher, 1991 Pos moll sp K/Pg US Atl CP 

Plank larva Jablonski & Hunt, 2006 NS moll sp Cret-Cz US CP 

Plank larva Jablonski, 1987 NS biv, gast sp Cret US CP 

Plank larva Stanley, 1986b NS biv sp Late Ng W N Amer 

Plank larva Valentine & Jablonski, 1986 NS biv, gast gen K/Pg US CP 

Schizodont Jablonski, 2008a Pos biv gen K/Pg Global May be example of trait hitch-hiking. Geographic range is more important 

in determining risk. 

Shell thick McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Sp richness Hoffman, 1986 NS biv gen K/Pg Euro 

Sp richness Jablonski, 1986 NS biv, gast sp K/Pg US CP 

Sp richness Jablonski, 1986 Neg biv, gast sp Late Cret US CP 

Sp richness Jablonski, 1989 NS biv, gast gen K/Pg N Am, Euro 

Sp richness Jablonski, 2005 NS biv, gast gen K/Pg US CP 

Sp richness Jablonski, 2005 Neg biv, gast gen Late Cret US CP 

Sp richness McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Sp richness Smith & Roy, 2006 Neg scallops gen Plio/Pleist California 

Sp richness Vermeij, 1986 NS moll sp Plio N Atl 

Stenohaline Hallam, 1981 Pos biv gen T/J Global 

Stenotherm McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Stenotherm Stanley, 1986a Pos biv sp Plio/Pleist SE US 

Stenotherm Stanley, 1990a Pos biv sp Late Ng W Atl 

Stenotopic Anderson & Roopnarine, 2003 NS corbulids gen Mio E Pacific Examined within phylogenetic framework. 

Stenotopic Hansen & others, 1993 NS moll sp K/Pg E Texas Used lithofacies tolerance as proxy for stenotopy. 

Stenotopic Kauffman, 1977 Pos biv sp Cret W Interior 

Stenotopic Kauffman, 1978 Mixed biv sp Cret N Amer 

Stenotopic Levinton, 1974 Pos 4 groups gen Phan Global Comparison of mortality rates between selective suspension-feeders 


23




level 

Stenotopic Posenato, 2009 Varies biv, brach gen Perm W Tethys Selectivity varies according to site. 

Strat range Stilwell, 2003 Neg moll gen, sp K/Pg S hemis 

Susp feeding Aberhan & others, 2007 Pos moll dom sp K/Pg Argentina 

Susp feeding Crampton & others, 2010 NS biv sp Cz NZ 

Susp feeding Hansen & others, 1987 Pos? biv sp K/Pg E Texas Shift from susp to dep feeders. May or may not be extinction selectivity. 

Susp feeding Hansen, Farrell, & Upshaw, 1993 Pos? moll sp K/Pg E Texas Shift from susp to dep feeders at one site. May or may not be extinction 

selectivity. 

Susp feeding Hansen & others, 1993 NS moll sp K/Pg E Texas Long-term shift from susp to dep feeders. Both go extinct at boundary. 

Susp feeding Hautmann & others, 2008 Pos biv dom sp T/J Tibet 

Susp feeding Jablonski & Raup, 1995 Pos biv gen K/Pg Global Selectivity due to low extinction in Nuculoida and Lucinoida. 

Susp feeding Jablonski, 1996a Pos biv gen K/Pg Global Argues that this is a taxonomic pattern. 

Susp feeding Kauffman, 1978 Pos biv sp Cret N Am 

Susp feeding Levinton, 1974 Pos 4 groups gen Phan Global Comparison of mortality rates between selective suspension-feeders 


Susp feeding McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Susp feeding Raup & Jablonski, 1993 Pos biv gen K/Pg Global Pattern due to low extinction in Nuculoida and Lucinoida. 

Susp feeding Rhodes & Thayer, 1991 Pos biv gen K/Pg Global 

Susp feeding Sheehan & Hansen, 1986 Pos moll sp K/Pg E Texas 

Susp feeding Stilwell, 2003 Pos moll gen, sp K/Pg S hemis 

Tax structure Rivadeneira & Marquet, 2007 NS biv sp Late Ng W S Amer 

Tropical Stanley, 1986a Pos biv sp Plio/Pleist SE US 

Water depth Jablonski & Raup, 1995 NS biv gen K/Pg US CP 

Water depth Jablonski, 1996a NS biv, gast gen K/Pg Global 

Water depth Levinton, 1973 Pos 6 species sp Rec New York Epifaunal taxa only. 

Water depth McClure & Bohonak, 1995 NS biv gen K/Pg US CP 

Water depth Raup & Jablonski, 1993 NS biv gen K/Pg Global 

Water depth Valentine & Jablonski, 1993 Pos moll sp Pleist California 

Key: 

Traits: Active locom = active locomotion; Aragonite = aragonitic shell composition; Bathy range = bathymetric range; Bur depth = burrowing depth; Escalation = escalated traits; Geo range = geographic range; 

Hypercapnia = sensitivity to hypercapnia; Meta rate = metabolic rate; Morph com = morphological complexity; Morph var = morphological variation; Multivariate = multivariate approaches; Plank larva 

= planktonic larva; Shell thick = shell thickness; Sp richness = species richness; Stenotherm = Stenothermal; Strat range = stratigraphic range; Susp feeding = suspension feeding; Tax structure = taxonomic 

structure. 

Pattern: Neg = negative selectivity (i.e., these taxa or taxa with higher levels of these traits are less likely to go extinct); NS = not significant; Pos = positive selectivity (i.e., these taxa or taxa with higher levels of 

these traits are more likely to go extinct); ? = unclear whether this pattern is due to extinction selectivity. 

Taxon: ammon = ammonites; biv = bivalve; brach = brachiopods; dom = dominated; gast = gastropods; moll = mollusks; superfm = superfamilies. 

Taxon level: fam = families; gen = genera; sp = species. 

Age: Cret = Cretaceous; Cz = Cenozoic; Eo = Eocene; E/O = end-Eocene extinction; F/F = late Devonian extinction; Jur = Jurassic; K/Pg = end-Cretaceous extinction; Mio = Miocene; Ng = Neogene; Nor = 

Norian; O/S = end-Ordovician; Perm = Permian; P/T = end-Permian extinction; Phan = Phanerozoic; Pleist = Pleistocene; Pliens = Pliensbachian; Plio = Pliocene; Pz = Paleozoic; Rec = Recent; Rhaet = 

Rhaetian; T/J = end-Triassic extinction; Toar = Toarcian; Tri = Triassic. 

Location: Am = America; Atl = Atlantic; Carib = Caribbean; CP = Coastal Plain; E = East; Euro = Europe; hemis = hemisphere; N = North; NZ = New Zealand; S = South; trop = tropical; W = West. 

Notes: dep = deposit; susp = suspension.

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